Quaternary International 443 (2017) 52e64 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint Sixty years after ‘The mastodonts of Brazil’: The state of the art of South American proboscideans (Proboscidea, Gomphotheriidae) * Dimila Mothe a, b, , Leonardo dos Santos Avilla a, c, Lidiane Asevedo a, d, Leon Borges-Silva a, Mariane Rosas e, Rafael Labarca-Encina f, Ricardo Souberlich g, Esteban Soibelzon h, i, Jose Luis Roman-Carrion j, Sergio D. Ríos k, Ascanio D. Rincon l, Gina Cardoso de Oliveira b, Renato Pereira Lopes m a Laboratorio de Mastozoologia, Departamento de Zoologia, Instituto de Bioci^encias, Universidade Federal do Estado do Rio de Janeiro, Av. Pasteur, 458, 501, Urca, CEP 22290-240, Rio de Janeiro, Brazil b Programa de Pos-graduaç ao~ em Geoci^encias, Centro de Tecnologia e Geoci^encias, Universidade Federal de Pernambuco, Rua Acad^emico Helio Ramos, s/n, Cidade Universitaria, CEP 50740-467, Recife, Brazil c Programa de Pos-graduaç ao~ em Biodiversidade Neotropical, Instituto de Bioci^encias, Universidade Federal do Estado do Rio de Janeiro, Av. Pasteur, 458, 501, Urca, CEP 22290-240, Rio de Janeiro, Brazil d Faculdade de Geoci^encias (Fageo), Campus Cuiaba, Universidade Federal de Mato Grosso, Av. Fernando Correa da Costa, 2367, Jardim Petropolis, CEP 78070-000, Cuiaba, Mato Grosso, Brazil e Laboratorio de Paleontologia, Centro de Ci^encias Agrarias, Ambientais e Biologicas, Universidade Federal do Reconcavo^ da Bahia, Cruz das Almas, Bahia, Brazil f Laboratorio de Paleoecología, Instituto de Ciencias Ambientales y Evolutivas, Universidad Austral de Chile, Casilla 567, Valdivia, Chile g Laboratorio de Paleontología, Departamento de Geología, Facultad de Ciencias Exactas y Naturales, Acceso Av. Mcal. Lopez, Universidad Nacional de Asuncion, Paraguay h Division Paleontología Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Paseo del Bosque s/n, 1900, La Plata, Argentina i CONICET, Argentina j Museo de Historia Natural “Gustavo Orces V.”, Instituto de Ci^encias Biologicas, Escuela Politecnica Nacional, Ladron de Guevara E11e253, Casilla Postal 17e01e2759, Quito, Ecuador k Departamento de Arqueología y Paleontología Secretaria Nacional de Cultura, Paraguay l Instituto Venezolano de Investigaciones Científicas, Laboratorio de Paleontologia, Centro de Ecología, Km 11 de la Carretera Panamericana, Apartado Postal 20632, Caracas 1020A, Venezuela m Programa de Pos-Graduaç ao~ em Geoci^encias, UFRGS, Av. Bento Gonçalves, 9500, CEP 91540-000, Porto Alegre, Rio Grande do Sul, Brazil article info abstract Article history: Studies on South American Gomphotheriidae started around 210 years ago and, 150 years later, the Available online 30 September 2016 classic study “The mastodonts of Brazil” by Simpson and Paula Couto (1957) attempted to clarify the complex issues related to our understanding of these proboscideans. Here, we update state of knowledge Keywords: regarding proboscideans in South America subsequent to the publication of Simpson and Paula Couto Notiomastodon (1957). The taxonomy of South American proboscideans is now stable and two species are recognized, Cuvieronius Notiomastodon platensis and Cuvieronius hyodon. The former had a wide distribution in South America Proboscidea ́ (from lowlands to highlands and from east to west coasts), while the latter was restricted to Ecuador, Extinction Paleoecology Peru and Bolivia. Although records of Notiomastodon are abundant and occur in almost overlapping Taxonomy geographic distribution with Cuvieronius, they have never been recorded in the same locality. Here, we evaluated over 500 South American localities with proboscidean remains, although only cranial and dental specimens show recognizable diagnostic features. As both proboscideans in South America had a generalist-opportunist alimentary strategy, a competitive exclusion probably precluded their sympatry. Their origin is most probably related to independent migrations from Central America during the Great * Corresponding author. Laboratorio de Mastozoologia, Departamento de Zoologia, Instituto de Biociencias,^ Universidade Federal do Estado do Rio de Janeiro, Av. Pasteur, 458, sala 501, Urca, CEP 22290-240, Rio de Janeiro, Brazil. E-mail addresses: [email protected] (D. Mothe), [email protected] (L. dos Santos Avilla), [email protected] (L. Asevedo), [email protected] (L. Borges-Silva), [email protected] (M. Rosas), [email protected] (R. Labarca-Encina), [email protected] (R. Souberlich), [email protected] (E. Soibelzon), [email protected] (J.L. Roman-Carrion), [email protected] (S.D. Ríos), [email protected] (A.D. Rincon), [email protected] (G.C. de Oliveira), [email protected] (R.P. Lopes). http://dx.doi.org/10.1016/j.quaint.2016.08.028 1040-6182/© 2016 Elsevier Ltd and INQUA. All rights reserved. D. Mothe et al. / Quaternary International 443 (2017) 52e64 53 American Biotic Interchange. They are not sister-taxa e Cuvieronius hyodon is sister-taxon of Rhyncho- therium, and this clade is closer to Notiomastodon platensis than to the other proboscideans, supporting the hypothesis of independent origins. Notiomastodon platensis has a continuous record from the Early Pleistocene to Early Holocene, when it became extinct, probably due to synergy of human impact and climatic changes during the Pleistocene/Holocene transition. In contrast, extinction of Cuvieronius hyodon happened much earlier, and it was not related to the terminal Pleistocene event that lead the extinction of selected megafauna in South America, including Notiomastodon. © 2016 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction Concepcion, Chile (Cuvier, 1806). Both specimens were described and figured by Cuvier (1806), who failed to assign generic or spe- After the publication of the first record of a South American cific names (binomial system of Linnaeus) in their descriptions and proboscidean 210 years ago (Cuvier, 1806), the taxonomic history of referred to them using the informal terms “mastodonte des this group had several chapters, and most researchers consider the cordilleres ” and “mastodonte humboldien”, respectively (Fig. 2). revision of Simpson and Paula Couto (1957) a watershed (Prado Fischer (1814) proposed the first valid specific name, Masto- et al., 2005; Shoshani and Tassy, 2005; Ferretti, 2010). Why is this therium hyodon, for the specimen from Ecuador (“mastodonte des classic paper so important? It comprises the first effort in 150 years cordilleres ”), and Cuvier (1824) later formally named these mate- to review the taxonomy and to congregate information on pro- rials as Mastodon andium (specimen from Ecuador) and Mastodon boscideans from many localities in South America; a contrast to humboldtii (specimen from Chile). However, the name Mastothe- previous traditional taxonomical studies that frequently proposed rium hyodon had priority over the name M. andium proposed by new names. The only exception is Cabrera (1929), a paper that was Cuvier (1824). Nevertheless, the isolated teeth described by Cuvier most certainly the basis of the study by Simpson and Paula Couto (1806, 1824) and Fischer (1814) are not diagnostic to any bunodont (1957). In the following six decades, due to more critical studies, proboscidean at the specific level, based on recent diagnoses, and the taxonomy of South American proboscideans further stabilized they could represent either of the South American taxa, including (Mothe et al., 2012; Mothe and Avilla, 2015). Inspired by Simpson the same taxon (Mastotherium hyodon and/or Mastodon humboldtii; and Paula Couto (1957), here we gathered a group of South Amer- Ficcarelli et al., 1995; Ferretti, 2008; Lucas, 2008). ican researchers to present the current state of the knowledge on Lund (1839) mentioned some proboscidean remains from Lagoa taxonomy, phylogeny, geographic distribution, chronology and the Santa, Minas Gerais State in Brazil, but he gave no classification, feeding paleoecology of South American proboscideans. description or illustration of the specimens (a nomen nudum), and Lesson (1842) listed only the name “Mastodon brasiliensis Lund”. 2. Materials and methods Almost seven decades later, when describing the specimens mentioned by Lund (1839), Winge (1906) referred to them as The information presented here includes a detailed survey of the “Mastodon andium”, although Mastotherium hyodon had priority literature of the last 210 years, since the first record of Proboscidea over this name for almost a century (Fig. 2). Furthermore, Winge in South America (Cuvier, 1806). We also comment and make new (1906) did not validate the name “Mastodon brasiliensis”. Only in inferences on taxonomy, geographic distribution, systematics and Hoffstetter (1952), when he properly published a systematic paleoecology based on more recently collected data. The specimens description of “Stegomastodon brasiliensis”, was the name credited of proboscideans from all over South America were, for the first to Lund and became valid. However, it was antedated by several time, catalogued in a single database (Supplementary Materials e other names proposed before his study (see Cabrera,1929; Simpson Table 1), reviewed and classified according to the current pro- and Paula Couto, 1957; Ferretti, 2008; Lucas, 2008; and Fig. 2). posed taxonomic classification (Mothe and Avilla, 2015). Each At the end of the 19th century and beginning of